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Foss. Rec., 19, 131–141, 2016 www.foss-rec.net/19/131/2016/ doi:10.5194/fr-19-131-2016 © Author(s) 2016. CC Attribution 3.0 License.

First occurrence of atrox (, ) in the Mexican state of Hidalgo and a review of the record of felids from the of Mexico

Victor Manuel Bravo-Cuevas1, Jaime Priego-Vargas2, Miguel Ángel Cabral-Perdomo1, and Marco Antonio Pineda Maldonado3 1Museo de Paleontología, Área Académica de Biología, Universidad Autónoma del Estado de Hidalgo, Ciudad del Conocimiento, Carretera Pachuca-Tulancingo km 4.5, CP 42184, Pachuca, Hidalgo, Mexico 2Área Académica de Biología, Doctorado en Biodiversidad y Conservación, Universidad Autónoma del Estado de Hidalgo, Ciudad del Conocimiento, Carretera Pachuca-Tulancingo km 4.5, CP 42184, Pachuca, Hidalgo, Mexico 3Área Académica de Biología, Maestría en Biodiversidad y Conservación, Universidad Autónoma del Estado de Hidalgo, Ciudad del Conocimiento, Carretera Pachuca-Tulancingo km 4.5, CP 42184, Pachuca, Hidalgo, Mexico

Correspondence to: Victor Manuel Bravo-Cuevas ([email protected])

Received: 26 May 2016 – Revised: 18 June 2016 – Accepted: 1 July 2016 – Published: 20 July 2016

Abstract. Panthera atrox was a common large-sized in the mesopredator level, such as the dire ( dirus ), during the . An isolated lower which has been also reported in the area. The high diversity canine and a fifth metacarpal bone referable to this of large herbivores recorded at southeastern Hidalgo, which were recovered from fluvial Quaternary deposits that outcrop in turn could represent potential prey of P. atrox, suggests in southeastern Hidalgo, central Mexico. Associated that some areas that now are part of central Mexico were suit- material belonging to Bison indicates a Rancholabrean North able sites for this large-sized cat. American Land Age; the age assignment is corrob- A review of the Pleistocene record of Felidae from Mex- orated by the presence of P. atrox. A comparative study with ico revealed that it encompasses 87.5 and 73.3 % of generic selected specimens of Panthera and indicates that and specific diversity known for North America, respec- the Hidalgoan sample shares the following diagnostic fea- tively, including seven genera (, Miracinonyx, Panthera, tures with P. atrox: a large, robust, and non-strongly curved , , , and Smilodon) and 11 species (Fe- lower canine; a large and relatively slender fifth metacarpal lis rexroadensis, Miracinonyx inexpectatus, Panthera atrox, with a well-developed projection on the palmar side at the Panthera onca, Puma concolor, Puma yagouaroundi, Lynx proximal end, narrow articulating surface for the unciform; rufus, Leopardus pardalis, Leopardus wiedii, Smilodon fa- a narrow notch on the articulating surface for the fourth talis, and Smilodon gracilis). The majority of these taxa have metacarpal; and a diaphysis that at the middle is oval in cross been reported from numerous late Pleistocene localities; in section. The record supplements the evidence of P. atrox in particular, Panthera atrox was relatively common and widely central Mexico and represents the first reported occurrence distributed across the Mexican . of this cat species in the state of Hidalgo. By the same token, the known geographic distribution of P. atrox in the Mexi- can territory suggests that it was relatively common in tem- perate areas of central Mexico between 19 and 24◦ N at an 1 Introduction altitudinal range from 1500 to 2250 m a.s.l. The large size (mean body mass of 300 kg) and hypercarnivorous adapta- The subfamily Pantherinae is a group of large-sized that tions of the American suggest it was the top predator originated in (probably in some part of what is now of the mammalian community recorded at southeastern Hi- ) during the end of the and at the beginning of dalgo, displacing other members of the guild at the Pliocene (10.8–3.8 Ma) (Johnson et al., 2006; Davis et al.,

Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin. 132 V. M. Bravo-Cuevas: First occurrence of Panthera atrox

2010; Tseng et al., 2016). This group of felids includes six 2 Study area extant species in the genera Panthera (P. leo (African lion), P. tigris (), P. onca (), and P. pardus ()), The material considered here has been recovered from a lo- ◦ 0 00 Uncia (U. uncia ()), and (N. nebulosa cality formally known as El Barrio (HGO-47: 20 07 41 N, ◦ 0 00 ()) (Davis et al., 2010; King and Wallace, 98 56 02.7 W) in the southeastern region of Hidalgo, cen- 2014); all of these species are considered to be vulnerable, tral Mexico (Fig. 1a). In this region Tertiary to Quaternary endangered, and/or threatened (IUCN, 2015). Extinct pan- volcanic and volcano sedimentary rock units crop out (IN- therines include the Pleistocene species P. gombaszoegensis EGI, 1992). The fossil-bearing strata consist of heteroge- (eastern ), P. fossilis (Eurasia), P. youngi (northeastern neous clastic material, including clay, silt, sand, and scarce China), P. spelaea (Eurasia and Alaska), and P. atrox (North conglomeratic lenses, setting in a fluvial depositional envi- America) (Kurtén and Anderson, 1980; Lange, 2002; Kurtén, ronment (Fig. 1b). 2009; Tseng et al., 2016). The oldest known pantherines are The associated mammalian fauna includes dire those referable to P. blytheae from the late Miocene–early (Canis dirus), giant armadillos (Glyptotherium floridanum), Pliocene of the Himalayan region (Tseng et al., 2016) and P. giant ground sloths (Paramylodon cf. harlani), llamas (Hemi- palaeosinensis from the Plio-Pleistocene of China (Mazák, auchenia gracilis and Camelops sp.), horses (Equus conver- 2010). sidens), pronghorns (Stockoceros conklingi and Capromeryx The late Pleistocene species P. atrox was a common large- minor), deer (Odocoileus cf. virginianus), bison (Bison sp.), sized cat in the North American subcontinent, which has gomphotheres (Cuvieronius sp.), and mammoths (Mam- been traditionally referred to as the (Kurtén muthus sp.) (Bravo-Cuevas, 2001, 2002; Bravo-Cuevas et al., and Anderson, 1980). Ancestors of the American lion appar- 2009, 2011, 2012, 2013). There is evidence of material be- ently crossed from Asia to Alaska sometime during the Illi- longing to the Bison, including a horn core fragment noian glacial stage (191 000 to 130 000 ago) (Lange, and an isolated tooth; the presence of this taxon is indica- 2002) at the beginning of the Rancholabrean North Ameri- tive of a Rancholabrean NALMA (Bell et al., 2004). The can Land Mammal Age (NALMA). Early representatives of biochronological age of the fossil assemblage is supported by P. atrox appeared during the Sangamonian interglacial stage, the presence of material belonging to Panthera atrox (present reaching areas of southern temperate North America in a rel- study). atively short time (ca. 125 000 years ago) (Kurtén and An- derson, 1980; Lange, 2002). In the Wisconsinan (75 000– 3 Materials and methods 10 000 years ago) at the end of the Rancholabrean NALMA, P. atrox spread across the United States and Mexico (Kurtén The sample consists of an isolated canine and a metacarpal and Anderson, 1980), and its southernmost occurrence has bone. The material is housed at the Sección de Macrover- been recorded in tropical areas of the Mexican territory tebrados, Museo de Paleontología, Universidad Autónoma (Montellano-Ballesteros and Carbot-Chanona, 2009; Carbot- del Estado de Hidalgo, México, with the catalogue numbers Chanona and Gómez-Pérez, 2014). UAHMP-4221 and UAHMP-4222, respectively. The phylogenetic status of Panthera atrox is controver- The anteroposterior and transverse diameters of the iso- sial, given that some authors have considered it to have been lated tooth were measured. The measurements of the closely related to (Simpson, 1941; Christiansen and metacarpal are those of Merriam and Stock (1932: table 62, Harris, 2009) and others that it was a subspecies of the ex- p. 134). Measurements in millimeters were taken using a dig- tant lion (Kurtén and Anderson, 1980). Recent studies on the ital caliper with a measuring range of 0–150 mm, a resolution phylogenetic relationship of P. atrox with other members of of 0.01 mm, and an accuracy of 0.003 mm. the genus Panthera indicate that it should be considered as a In to assess the taxonomic identity of the dental and separate species closely related to (P. leo) and (P. postcranial remains from Hidalgo, we performed a compara- tigris) (Barnett et al., 2009; Christiansen and Harris, 2009; tive study with selected specimens of Panthera atrox housed King and Wallace, 2014). at the George C. Page Museum, La Brea Discoveries, Mu- Information regarding the fossil material, description, and seum of Natural History of Los Angeles County (LACM), formal taxonomic characterization of Mexican specimens California, USA. Additional comparisons were done with referable to P. atrox is unknown in many instances. Hence, specimens belonging to the extant species P. leo (OCMP- the purpose of this study is to formally describe the dental 080) and P. tigris (OCMP-077) housed at the Osteologi- and postcranial material belonging to Panthera atrox from cal Collection of the Museo de Paleontología, Universidad the late Pleistocene of southeastern Hidalgo, central Mexico, Autónoma del Estado de Hidalgo. Furthermore, the Hidal- as well as to discuss some ecological and geographic aspects goan material was compared with specimens described and of the record. Furthermore, a review of the record of felids figured in Merriam and Stock (1932), Whitmore and Fos- from the Pleistocene of Mexico is provided. ter (1967), and Montellano-Ballesteros and Carbot-Chanona (2009).

Foss. Rec., 19, 131–141, 2016 www.foss-rec.net/19/131/2016/ V. M. Bravo-Cuevas: First occurrence of Panthera atrox 133

Figure 1. (a) Index map showing the study area in southeastern Hidalgo, central Mexico; the capital of the state (Pachuca) and the late Pleistocene locality El Barrio (HGO-47) are depicted. (b) Stratigraphic section of the El Barrio locality (HGO-47); the arrow indicates the fossil-bearing level.

In order to review the Pleistocene record of Felidae 4.1 Description from Mexico, we performed a bibliographic search in pri- mary sources (journal articles, books, and book chapters), The canine UAHMP-4221 is large and robust (Table 1). A secondary sources (abstracts, catalogs, and theses), and thin layer of enamel (< 1 mm thick) covers the first third databases (The Paleobiology Database (https://paleobiodb. of the tooth. The tip has broken anterolaterally after death org) and the Mexican Quaternary Database; (given that it does not show wear) and is subacute in shape. Arroyo-Cabrales et al., 2005). The collected data were ar- The specimen is moderately curved, and in cross section it is ranged in terms of its generic and specific diversity, repre- oval to oval-elongate toward to the base. The lateral surface senting the basis of a distribution map of the known (so far) of the tooth is slightly convex, whereas the medial surface is Pleistocene felids from the Mexican territory. flattened (Fig. 2). The metacarpal UAHMP-4222 is slender and long, show- ing a relatively gracile appearance (Table 2, Fig. 3). The shaft 4 Systematic paleontology is oval in cross section and without sharply defined borders between the dorsal and palmar surfaces. At the proximal end, – Order Bowdich, 1821 the articular surface for the unciform is narrow. The medial – Felidae Fischer, 1817 face presents the articular surface for metacarpal IV, formed by a large projection (“ear-shaped projection” of Merriam – Subfamily Pantherinae Pockock, 1917 and Stock, 1932) and a narrow notch placed on the palmar – Genus Panthera Oken, 1816 side. The lateral surface presents a convex tuberosity. At the distal end, the lateral tuberosity is more prominent than the – Panthera atrox Leidy, 1853 medial tuberosity. The distal articulation surface is situated at Referred material. El Barrio locality: UAHMP-4221, left a 10◦ angle from the axis of the shaft and a prominent palmar lower canine; UAHMP-4222, left fifth metacarpal. keel is apparent.

Distribution and age. The American lion was widely dis- 4.2 Taxonomic assessment tributed across North America from Alaska (Whitmore and Foster, 1967) to southern Mexico (Montellano-Ballesteros The deciduous lower canine of felids is distinguished by hav- and Carbot-Chanona, 2009; Carbot-Chanona and Gómez- ing an accessory cusp situated on the anterolingual side of the Pérez, 2014); some records from Alaska and western Canada tooth (Salles, 1992). The specimen UAHMP-4221 does not maybe represent members that are more closely related to the show this condition, indicating that it is a permanent tooth; cave lion (P. spelaea) (Barnett et al., 2009). It is a taxon lim- furthermore, it resembles the lower canine of pantherines ited to the Rancholabrean NALMA (Kurten and Anderson, in the absence of a lingual cavity (a widespread condition 1980; Lange, 2002). among felids), and differs from an upper canine in the ab- www.foss-rec.net/19/131/2016/ Foss. Rec., 19, 131–141, 2016 134 V. M. Bravo-Cuevas: First occurrence of Panthera atrox

Table 1. Comparison of measurements (in mm) in UAHMP-4221 and lower canine specimens of Panthera atrox (a–d) and the extant species P. tigris (e) and P. leo (f). Rancholabrean localities: (a) El Barrio, southeastern Hidalgo, central Mexico (present study); (b) Lost Chicken Creek, Alaska, United States (Whitmore and Foster, 1967: table 1, p. 250); (c) Rancho La Brea, California, United States (Merriam and Stock, 1932: table 94, p. 177); and (d) La Tejería, Chiapas, southern Mexico (Montellano-Ballesteros and Carbot-Chanona, 2009: table 1, p. 219). Abbreviations: AD, anteroposterior diameter; TD, transverse diameter.

(a) (b) (c) (d) (e) (f) UAHMP-4221 USNM-23619 Rancho La Brea IHNFG-2678 OCMP-077 OCMP-080 Left Right (N= 11) Right Left Right/left AD 30.1 30.0 21.8–30.4 21.9 20.8 21.0/20.7 TD 21.6 21.5 15.1–21.6 15.6 14.2 13.9/13.7

Table 2. Comparison of measurements (in mm) between UAHMP-4222 and fifth metacarpal specimens of Panthera atrox from Rancho La Brea (RLB), late Pleistocene of California, United States (Merriam and Stock, 1932: table 62, p. 134). The observed range in the specimens from Rancho La Brea is in parentheses.

UAHMP-4222 RLB RLB Left Right Left Greatest length 105.9 x = 105 x = 106.6 (90.6–115.3) (91.7–113.0) Greatest transverse diameter of proximal end 28.4 28.7 x = 27.8 (25.1–27.7) (24.9–30.0) Greatest dorsoventral diameter of proximal end 26.8 x = 31.3 x = 29.8 (27.0–30.8) (27.3–33.2) Transverse diameter at middle of shaft 17.0 x = 17.0 x = 16.3 (14.2–15.8) (14.2–17.6) Dorsoventral diameter at middle of shaft 13.9 x = 15.0 x = 14.1 (13.2–15.2) (12.2–15.8) Greatest transverse diameter at distal end of shaft 23.5 x = 24.9 x = 24.8 (21.2–25.2) (22.2–25.2) sence of a lingual ridge (Salles, 1992) and in being more ro- bust. By contrast, the lower canine of Smilodon is smaller and slender, strongly curved, and shows a median lateral ridge (Merriam and Stock, 1932). The size of the tooth is comparable to that observed for lower canines belonging to Panthera atrox, including USNM 23619 (right lower jaw with c, p3–m1) from Lost Chicken Creek, Rancholabrean of Fairbanks, Alaska; it falls in the upper limit of the observed range in specimens from Ran- cho La Brea, California, United States; and it is nearly 25 % larger than the specimen IHNFG-2678 (isolated lower ca- nine) from Chiapas, southern Mexico (Table 1). The differ- ence in size between the specimens from Hidalgo and Chia- pas could be explained by intraspecific variation (related to age and/or sex), considering that UAHMP-4221 shows di- mensions comparable to the larger lower canines from Ran- cho La Brea, whereas the dimensions of IHNFG-2678 cor- respond to those of the smaller ones (Table 1). It should be stated that the specimen UAHMP-4221 is significantly larger Figure 2. Isolated left lower canine (UAHMP-4221) of Panthera (ca. 33 %) than lower canines of Panthera leo and P. tigris; atrox from the late Pleistocene of southeastern Hidalgo, central (a) (b) however, the size of this tooth between those extant species Mexico. Labial and lingual views. Scale bar equals 2 cm. is similar (Table 1).

Foss. Rec., 19, 131–141, 2016 www.foss-rec.net/19/131/2016/ V. M. Bravo-Cuevas: First occurrence of Panthera atrox 135

Figure 3. Left fifth metacarpal (UAHMP-4222) of Panthera atrox from the late Pleistocene of southeastern Hidalgo, central Mexico. (a) Ventral, (b) dorsal, (c) medial, (d) lateral, (e) proximal, and (f) distal views. Scale bar equals 2 cm.

Among felids, the morphology of the limb elements is ciform, and a relatively stout diaphysis (Merriam and Stock, somewhat homogeneous, including the metacarpals. In gen- 1932). eral, the metacarpal bones of felids are characterized by be- The comparative study indicates that the size and mor- ing short and robust with a broad and curved diaphysis as phology of dental and postcranial remains from Hidalgo are well as broad proximal and narrow distal ends (Morales- closely comparable to those observed in specimens belong- Mejía and Arroyo-Cabrales, 2012). All these features are ob- ing to Panthera atrox. Hence, the studied sample is formally served in the specimen UAHMP-4222, indicating its felid assigned to that large-sized cat species. condition. In particular, the fifth metacarpal from Hidalgo resembles those of Panthera atrox in the following charac- ters: (1) well-developed projection on the palmar side at the 5 Paleobiological significance proximal end; (2) the articulating surface for the unciform is 5.1 Paleoecology narrow; (3) the diaphysis is relatively slender; (4) the notch on the articulating surface for the fourth metacarpal is nar- The American lion was one of the largest cats that inhabited row; and (5) the diaphysis at the middle is oval in cross sec- North America during the late Pleistocene (Kurtén and An- tion (Merriam and Stock, 1932). Furthermore, the size of derson, 1980; Lange, 2002). Body size estimations indicate = UAHMP-4222 (greatest length 105.9 mm) is within the that this felid had a mean body mass of 300 kg, ranging from observed range of fifth metacarpals of P. atrox from Rancho 200 to 400 kg (Van Valkenburgh et al., 2016). It has been ob- La Brea, California, United States (Table 2); however, it is served that prey body size tends to increase with the predator about 25 % larger than the fifth metacarpal of OCMP-077 be- size (Sinclair et al., 2003). Therefore, it should be expected = longing to P. tigris (greatest length 80.3 mm). It should be that large mammalian herbivores were common prey of Pan- noted that the fifth metacarpal of Smilodon is distinguished thera atrox. Van Valkenburgh et al. (2016: fig. 2, p. 865) pre- in having a poorly developed projection on the palmar side dicted a typical prey size ranging from 60 to 900 kg for the at the proximal end, a broad articulating surface for the un- American lion, and the maximum prey size could have been about 1000 kg. www.foss-rec.net/19/131/2016/ Foss. Rec., 19, 131–141, 2016 136 V. M. Bravo-Cuevas: First occurrence of Panthera atrox

Figure 4. Potential common prey-size range for Panthera atrox from the late Pleistocene of southeastern Hidalgo, including the herbivores that have been reported at the El Barrio locality (HGO-47). Diamond and line indicate the mean and observed range of body mass (from Van Valkenburgh et al., 2016).

Given the above and considering the taxonomic mam- ) suggest that this extinct cat occupied the top malian composition known at the El Barrio locality, it seems of the trophic chain, considering that both conditions are typ- that potential prey for Panthera atrox in that site could in- ical of extant top predators (Ritchie and Johnson, 2009). At clude adult individuals of llamas (Hemiauchenia gracilis the El Barrio locality, it is probable that the individual of P. and Camelops sp.), deer (Odocoileus cf. virginianus), horses atrox described here had the ecological role of the top preda- (Equus conversidens), and even bison (Bison sp.); the body tor, displacing other of small to medium size, such mass among these herbivores is estimated to have been 200 as the dire wolf (Canis dirus) to the mesopredator guild. to 900 kg (Fig. 4). Hunting in groups increases the upper range of available prey size; consequently, it has been sug- 5.2 Geographic distribution gested that large-sized Pleistocene cats (such as Panthera atrox) were able to kill prey with a body mass of about By the late Pleistocene, the American lion was widely dis- 6000 kg (Van Valkenburgh et al., 2016). Assuming a hunting tributed from Alaska to southern Mexico. The earliest known group behavior and potential presence of other individuals occurrences are from the Sangamonian interglacial stage, in- belonging to Panthera atrox at southeastern Hidalgo during cluding localities in the western of the United States, as well the late Pleistocene, it is also probable that adult individu- as northern and central Mexico (Kurtén and Anderson, 1980; als of glyptodonts (Glyptotherium floridanum) and ground Van Devender et al., 1985), indicating that this large-sized sloths (Paramylodon cf. harlani), as well as young and/or cat reached regions of southern temperate North America in a subadult individuals of proboscideans (Cuvieronius sp. and relatively short time. Subsequently, the American lion spread Mammuthus sp.), could represent other, perhaps occasional its distribution to the Great Plains, the Great Basin, the Cal- prey. ifornia Coast, the Gulf Coast, and Mexico during the Wis- The American lion was the second largest carnivore in the consinan (Kurtén and Anderson, 1980; Lange, 2002). Based late Pleistocene ecosystems of North America, exceeded by on the known geographic distribution of Panthera atrox, it only the short-faced Arctodus simus (Kurtén and Ander- seems that it was a common inhabitant of temperate areas of son, 1980). The large size and restricted dietary behavior of central-western North America, although it was able to reach Panthera atrox (a carnivore that feeds mostly on meat, i.e., a tropical areas that now are part of southern Mexico (Fig. 5).

Foss. Rec., 19, 131–141, 2016 www.foss-rec.net/19/131/2016/ V. M. Bravo-Cuevas: First occurrence of Panthera atrox 137

Figure 5. Geographic distribution of Panthera atrox in North Amer- ica during the late Pleistocene (main source: Kurtén and Anderson, 1980). The gray silhouette indicates the record from southeastern Hidalgo, central Mexico.

Previous to this study, the American lion Panthera atrox has been reported from nine Mexican localities in northern (La Brisca (Sonora); Arroyo-Cabrales et al., 2005), central (San Josecito (Nuevo León), El Cedral (San Luis Potosí), Figure 6. Mexican Pleistocene localities with records of Panthera El Cedazo (Aguascalientes), Chapala–Zacoalco (Jalisco), atrox, considering (a) a hypsometric base and (b) the Mexican bio- Tequixquiac (State of Mexico); Freudenberg, 1910; Mooser geographic corridors (sensu Ceballos et al., 2010). Diamond indi- and Dalquest, 1975; Lorenzo and Mirambell, 1981; Arroyo- cates the record from southeastern Hidalgo, central Mexico. Bio- Cabrales and Polaco, 2003; Arroyo-Cabrales et al., 2005; Lu- geographic corridors: 1, eastern US–Sierra Madre Oriental; 2, west- cas, 2008), and southern (La Simpatía, La Tejería y Villa ern US–Baja California; 3, Rocky Mountains–Sierra Madre Oc- Corzo (Chiapas); Aviña, 1969; Montellano-Ballesteros and cidental; 4, central US–northern Mexico; 5, Transvolcanic Belt– Carbot-Chanona, 2009; Carbot-Chanona and Gómez-Pérez, Sierra Madre del Sur; 6, Tamaulipas–Central America Gulf Low- 2014) areas of the country. The record reported here sup- lands; and 7, Sonora–Central America Pacific Lowlands. plements its presence in central Mexico, specifically in ar- eas that now are part of southeastern Hidalgo. It is noted that Sierra Madre Occidental branch and to the Great Plains– most of the localities where Panthera atrox has been reported western Chihuahua/Central Plateau corridors, respectively are located in central Mexico, between 19 and 24◦ N at an al- (after Ferrusquía-Villafranca et al., 2010). Furthermore, it titude from 1500 to 2250 m a.s.l. (including El Barrio locality used the corridor of the Tamaulipas – Central America Gulf at 2184 m a.s.l., Hidalgo (present study)), whereas the occur- Lowlands (i.e., Gulf Coastal Plain corridor (after Ferrusquía- rences in northern (La Brisca, Sonora) and southern (Chi- Villafranca et al., 2010)) for reaching southern tropical areas apasan localities) Mexico are located at a mean altitude of of the Mexican territory (Fig. 6b). 750 m a.s.l. (Fig. 6a). According to the major biogeographic corridors of Ce- ballos et al. (2010), the records of Panthera atrox from 6 Review of the felid record from the Pleistocene of northwestern and central Mexico (including the Hidal- Mexico goan record considered in the present study) indicate that this felid used the corridors of the Rocky Mountains– In Mexico, Pleistocene felid are uncommon and Sierra Madre Occidental and/or central US–northern Mex- fragmentary. Nevertheless, this group of carnivores was ico, which correspond to the southern Rocky Mountains– relatively diverse, including seven genera (Felis, Miraci- www.foss-rec.net/19/131/2016/ Foss. Rec., 19, 131–141, 2016 138 V. M. Bravo-Cuevas: First occurrence of Panthera atrox

Table 3. The record of felids from the Pleistocene of Mexico. The morphotectonic provinces are those of Ferrusquía-Villafranca (1993). Morphotectonic provinces: NW, Northwestern Plains and Sierras; CH-CO, Chihuahua–Coahuila Plateaus and Ranges; SMOr, Sierra Madre Oriental; CeP, Central Plateau; TMVB, Trans-Mexican Volcanic Belt; GCP, Gulf Coastal Plain; SMS, Sierra Madre del Sur; CHI, Sierra Madre de Chiapas; YPL, Yucatan Platform. Localities: 1, El Golfo (Sonora); 2, La Brisca (Sonora); 3, Terapa (Sonora); 4, Cuatro Ciénegas (Coahuila); 5, Cueva de Jiménez (Chihuahua); 6, Cueva de San Josecito (Nuevo León); 7, El Cedral (San Luis Potosí); 8, Mina San Antonio (San Luis Potosí); 9, El Cedazo (Aguascalientes); 10, Chapala–Zacoalco (Jalisco); 11, Tequixquiac (State of Mexico); 12, Tlapacoya (State of Mexico); 13, Tlailotlacan (State of Mexico); 14, El Barrio (Hidalgo, present study); 15, Valsequillo (Puebla); 16, Mixtequilla (Veracruz); 17, San Agustín (Oaxaca); 18, La Simpatía (Chiapas); 19, La Tejería (Chiapas); 20, Villa Corzo (Chiapas); 21, Cueva de Loltún (Yucatán); 22, Hoyo Negro (Quintana Roo).

NW CH-CO SMOr CeP TMVB GCP SMS CHI YPL Taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Panthera atrox X XX X XXX XXX Panthera onca X X XX X X Panthera cf. P. onca X Puma concolor XX X XX X X X Puma yagouaroundi X X Lynx rufus X XX XX X XX Leopardus pardalis X Leopardus wiedii X Smilodon fatalis X X XX X Smilodon sp. X Felis rexroadensis X Miracinonyx inexpectatus X nonyx, Panthera, Puma, Lynx, Leopardus, and Smilodon) fatalis; Mooser, 1959; Mooser and Dalquest, 1975), Trans- and 11 species (Felis rexroadensis, Miracinonyx inexpecta- Mexican Volcanic Belt (P. atrox, P. onca, P. concolor, and S. tus, Panthera atrox, Panthera onca, Puma concolor, Puma fatalis; Freudenberg, 1910; Aviña, 1969; Rufolo, 1998; Lu- yagouaroundi, Lynx rufus, Leopardus pardalis, Leopardus cas, 2008; Ferrusquía-Villafranca et al., 2010), Gulf Coastal wiedii, Smilodon fatalis, and Smilodon gracilis) (Table 3). Plain (P. concolor; Arroyo-Cabrales et al., 2005; Ferrusquía- The Mexican record of Pleistocene felids includes 87.5 % Villafranca et al., 2010), Sierra Madre del Sur (P. con- and 73.3 % of generic and specific diversity known for North color; Arroyo-Cabrales et al., 2005; Ferrusquía-Villafranca America, respectively. et al., 2010), Sierra Madre de Chiapas (P. atrox; Montellano- As far as we know, the only record of felids from the Ballesteros and Carbot-Chanona, 2009; Carbot-Chanona and early Pleistocene of Mexico consists of three species (Fe- Gómez-Pérez, 2014), and Yucatan Platform (P. concolor and lis rexroadensis, Miracinonyx inexpectatus, and Panthera cf. S. fatalis; Álvarez and Polaco, 1982; Collins et al., 2015). P. onca) from El Golfo, State of Sonora, in the Northwest- The present study adds an occurrence of Panthera atrox from ern Plains and Sierras morphotectonic province (Lindsay, a late Pleistocene locality in southeastern Hidalgo, central 1984; Croxen III et al., 2007). By contrast, the late Pleis- Mexico, within the Trans-Mexican Volcanic Belt morphotec- tocene record of Mexican felids is represented by at least tonic province. eight species whose material has been recovered from sev- The species Puma yagouaroundi (jaguarondi) is known eral localities across the country (Fig. 7). from the Sierra Madre Oriental and Yucatan Platform mor- Fossil material that has been referred to Panthera atrox photectonic provinces (Arroyo-Cabrales and Johnson, 1998; (the American lion), P. onca (jaguar), Lynx rufus (bob- Arroyo-Cabrales and Álvarez, 2003). By the same token, the cat), Puma concolor (), and Smilodon fatalis (saber- species Leopardus pardalis () and a saber-tooth cat toothed cat) is somewhat numerous. These species are known with an uncertain specific identity (referred to as Smilodon from several late Pleistocene localities in the following mor- cf. S. gracilis) have single occurrences from localities in photectonic provinces: Northwestern Plains and Sierras (P. the Yucatan Platform and Trans-Mexican Volcanic Belt mor- atrox and P. onca; Arroyo-Cabrales et al., 2005; Ferrusquía- photectonic provinces, respectively (Kurtén, 1967; Arroyo- Villafranca et al., 2010), Chihuahua–Coahuila Plateaus and Cabrales and Álvarez, 2003; Ferrusquía-Villafranca et al., Ranges (P. onca, L. rufus, and P. concolor; Gilmore, 1947; 2010). Messing, 1986), Sierra Madre Oriental (P. atrox, P. onca, There is a report of Leopardus wiedii from a site known as L. rufus, P. concolor, and S. fatalis; Lorenzo and Miram- Tlailotlacan, late Pleistocene of the State of Mexico (Arroyo- bell, 1981; Arroyo-Cabrales and Álvarez, 2003; Arroyo- Cabrales et al., 2005, 2010). Furthermore, there are mentions Cabrales et al., 2005, 2010; Ferrusquía-Villafranca et al., of Smilodon sp., Panthera sp., and Lynx sp. from localities in 2010), Central Plateau (P. atrox, P. onca, L. rufus, and S. the Sierra Madre Oriental (Smilodon sp.; Arroyo-Cabrales et

Foss. Rec., 19, 131–141, 2016 www.foss-rec.net/19/131/2016/ V. M. Bravo-Cuevas: First occurrence of Panthera atrox 139

Figure 7. Mexican Pleistocene localities with records of felids. The map is regionalized in the morphotectonic provinces of Ferrusquía- Villafranca (1993). Abbreviations of the morphotectonic provinces as in Table 3. Squares indicate the early Pleistocene localities and circles the late Pleistocene localities. The felid record includes the following species: A: Panthera atrox; B: Panthera onca; C: Puma concolor; D: Puma yagouaroundi; E: Lynx rufus; F: Leopardus pardalis; G: Leopardus wiedii; H: Smilodon fatalis; I: Smilodon cf. S. gracilis; J: Felis rexroadensis; K: Miracinonyx inexpectatus. al., 2005) and Trans-Mexican Volcanic Belt morphotectonic agnostic features: a large, robust, and non-strongly curved provinces (Smilodon sp., Panthera sp., and Lynx sp.; Arroyo- lower canine; a large and relatively slender fifth metacarpal Cabrales et al., 2005; Bravo-Cuevas et al., 2009). with a well-developed projection on the palmar side at the Clearly, felids were more diverse and widespread in their proximal end; a narrow articulating surface for the unci- geographic distribution across the Mexican territory during form; a narrow notch on the articulating surface for the fourth the late Pleistocene. There are several areas in northern, cen- metacarpal; and a diaphysis that at the middle is oval in cross tral, and southern Mexico that testify to the presence of section. The recovery of the American lion at the El Barrio three to four species of felids, including Cuatro Ciénegas, locality suggests that this large-sized hypercarnivore was the Coahuila (P. onca, P. concolor, and L. rufus); El Cedazo, top predator of the mammalian communities that inhabited Aguascalientes (P. atrox, P. onca, L. rufus, and S. fatalis); southeastern Hidalgo during the late Pleistocene. In addition, Chapala–Zacoalco, Jalisco (P. atrox, P. onca, P. concolor, it seems that some areas that now are part of central Mexico and S. fatalis), Tequixquiac, State of Mexico (P. atrox, L. ru- were suitable hunting sites for members of P. atrox, consider- fus, and S. fatalis); and Cueva de Loltún, Yucatán, (P. con- ing the high diversity of large mammalian herbivores (body color, P. yagouaroundi, and L. pardalis). The San Josecito mass ≥ 200 kg) that have been recorded there. In addition, Cave locality testifies to the presence of six species of felids. Panthera atrox was relatively common in temperate areas of It is probable that the high diversity recorded there should central Mexico between 19 and 24◦ N. be related to the nature of the site, which functioned as a The Pleistocene record of Felidae from Mexico encom- shelter for maternity, resting, and/or feeding of several carni- passes 87.5 and 73.3 % of total generic and specific felid di- vore groups, including felids (Arroyo-Cabrales and Álvarez, versity, respectively, known for North America. 2003).

Acknowledgements. We are indebted to Blaire Van Valkenburgh and Ismael Ferrusquía Villafranca for their comments and sugges- 7 Conclusions tions, which greatly improved the final version of the manuscript. The first author thanks Aisling Farrell, collection manager of An isolated lower canine and a fifth metacarpal bone recov- the Page Museum, La Brea Tar Pits, for her kind support during ered from fluvial Quaternary deposits in southeastern Hi- his visit to that collection. Thanks to Florian Witzmann and dalgo, central Mexico, are formally described and assigned Anna Wenzel for their help along the editing and handling process to Panthera atrox. A comparative study indicates that the Hi- of the manuscript. Last but not least, thanks to the Museum für dalgoan specimens share with that species the following di- Naturkunde in Berlin for defraying the article-processing charges. www.foss-rec.net/19/131/2016/ Foss. Rec., 19, 131–141, 2016 140 V. M. Bravo-Cuevas: First occurrence of Panthera atrox

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